Xylem anatomical and hydraulic traits vary within crown but not respond to water and nitrogen addition in Populus tomentosa

Abstract

Nitrogen (N) and water availability are the two crucial factors confining tree growth and forest productivity. Irrigation, fertilization, and combined fertigation are commonly applied to plantation forests for improving productivity. However, how xylem vasculature responds to these management practices remains poorly understood. Here we investigated the responses of tree growth, xylem anatomical structure, and hydraulics of the upper, middle, and lower canopy branches in a 6-year-old Populus tomentosa plantation subjected to four years of irrigation and fertigation treatments. The results showed that an improvement in tree growth only occurred in the irrigation treatment while the addition of N in fertigation treatments did not have a cumulative effect on tree growth. Most of the xylem anatomical traits, including vessel hydraulic diameter (Dh), vessel density (VD), vessel fraction (VF), double vessel wall thickness (t), potential specific sapwood hydraulic conductivity (Kp), fiber wall to lumen ratio (Tf/Df), vessel wall reinforcement (t/b)2, and wood density (WD) all showed conserved plasticity to long-term irrigation and fertigation, despite different tree growth rate, implying that aboveground biomass accumulation may have decoupled from the branch-level xylem traits. Besides, the alterations of other factors, including stomatal regulation strategy and crown structure with soil water and N availabilities, might better explain the variation in tree growth. In contrast, greater variations in branch xylem traits were detected across canopy layers. The upper canopy showed greater hydraulic safety characterized by narrower and denser vessels, higher cell wall to lumen ratio, and higher wood density compared to the lower canopy, while hydraulic efficiency (Kp) remained constant across the crown, thus highlighting the priority of hydraulic safety over efficiency in the construction of branch xylem. Overall, our study revealed the response pattern of P. tomentosa xylem structure to long-term water and N management, which facilitates a comprehensive understanding on the mechanisms underpinning the influence of water and nutrients regulation on the performance of fast-growing tree species’ plantations.